A Thesis submitted to the Faculty of Graduate Studies and Research in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Biology, University of Regina, xvi, 199 p. : maps

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The ability of predator populations to expand their ranges and adapt to new environments is often attributed to having a generalist dietary strategy, which is thought to be represented both at the population and individual level. Cormorants (Phalacrocorax spp.) are considered to be opportunistic generalists capable of using a wide variety of aquatic prey. This reputation is partially responsible for the global conflict between piscivorous cormorants and fish harvesters, which is one of the most widespread wildlife management issues in history. Despite the persistent belief that cormorants adversely affect economically important fish populations, relatively little is known about their trophic ecology and habitat use. Stable nitrogen and carbon isotopes are popular tools for studying food webs, and offer a comprehensive assessment of diet, trophic position, and ecological niche when combined with traditional diet analyses. However, the interpretation of isotope data may be confounded by variation in the lipid content of sample tissues. No validated lipid-normalization procedures are currently available for any cormorant species, or any fish-eating birds. As such, I first determined the effect of lipids on the stable carbon and nitrogen isotopes (δ13C and δ15N) values in cormorant tissues, and tested three published lipid-normalization models on stable isotope signatures in double-crested cormorant (P. auritus) muscle and liver tissues. The presence of lipids in cormorant muscle and liver altered the stable isotopes values, indicating corrections were required. However, the effects of lipids in cormorants were unpredictable and thus violated a major assumption of published lipid-normalization models. As a result, lipids must be chemically removed from cormorant muscle and liver tissue. I then examined the diet and trophic position of breeding populations of double-crested cormorants from three different lakes. The results revealed that cormorants generally occupied top-predator positions and relied heavily on pelagic prey in all food webs examined. The isotopic values of cormorants and pelagic predatory fish were sometimes similar, suggesting that dietary overlap is possible. To determine whether cormorants are true dietary generalists I studied double-crested cormorants from breeding colonies spanning three major ecoregions. Analyses of stomach contents revealed that at the population level cormorant diet varied widely by location, likely reflecting local food-web structure. However, within populations individuals were much more specialized than expected. Temporal shifts in δ13C and δ15N values in cormorant tissues with different turnover rates (muscle vs. liver) indicated that foraging varied among populations. The dietary niche occupied by cormorants will affect their interactions with fish, highlighting the importance of understanding their impacts to fish populations both at the population and individual level. Ultimately, my research has shown that cormorants do not consume prey indiscriminately, and instead may have more specific and uniform dietary niche requirements than previously considered. From a management perspective, cormorants should not be assumed to have negative effects on fish in all situations; however, further attention is required to determine the impacts of dietary overlap with sport fish. Ecologically, I have shown that generalist species can be much more consistent and specialized than previously considered. Further, individuals within generalist species may be highly specialized, which will change the overall effects of the population on other species in the food web.

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A Thesis Submitted to the Faculty of Graduate Studies and Research In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy *, University of Regina. *, * p.